Response of the whitespotted sawyer beetle, Monochamus s. scutellatus, and associated woodborers to pheromones of some Ips and Dendroctonus bark beetles

The response of whitespotted sawyer beetle, Monochamus s. scutellatus, to pheromones of the bark beetles, Dendroctonus rufipennis, Ips pini, Ips perturbatus and Ips latidens, and α‐pinene was investigated with field‐trapping experiments. Traps baited with ipsenol caught significantly more M. s. scutellatus than unbaited traps, whereas the other compounds (ipsdienol, ipsdienol plus lanierone, ipsdienol plus cis‐verbenol or frontalin) did not. Combining α‐pinene with ipsdienol, ipsdienol plus lanierone, ipsdienol plus cis‐verbenol or with frontalin did not increase captures of M. s. scutellatus above those of α‐pinene alone, whereas the combination of α‐pinene with ipsenol did. When α‐pinene was combined with ipsdienol or frontalin, trap captures of Monochamus mutator were significantly higher than unbaited traps or traps baited with frontalin but were not higher than traps baited with α‐pinene. The combination of ipsenol and α‐pinene was significantly more attractive to Monochamus notatus than unbaited traps; however, traps containing either ipsenol or α‐pinene were as attractive as the combination. None of the species of Buprestidae (Buprestis maculativentris and Chalcophora virginiensis) responded significantly to any of the treatments.     

Selective manipulation of predators using pheromones: responses to frontalin and ipsdienol pheromone components of bark beetles in the Great Lakes region

Abstract 1 One proposed approach to improving biological control of bark beetles (Coleoptera: Scolytidae; alt. Curculionidae: Scolytinae) is to manipulate predator movement using semiochemicals. However, selective manipulation is impeded by attraction of both predators and pests to bark beetle pheromones. 2 The primary bark beetle affecting pine plantations in Wisconsin, U.S.A., is the pine engraver, Ips pini (Say). Other herbivores include Ips grandicollis (Eichhoff) and Dryophthorus americanus Bedel (Curculionidae). The predominant predators are the beetles Thanasimus dubius (Cleridae) and Platysoma cylindrica (Histeridae). 3 We conducted field assays using two enantiomeric ratios of ipsdienol, and frontalin plus α‐pinene. Ipsdienol is the principal pheromone component of I. pini, and frontalin is produced by a number of Dendroctonus species. α‐Pinene is a host monoterpene commonly incorporated into commercial frontalin lures. 4 Thanasimus dubius was attracted to frontalin plus α‐pinene, and also to racemic ipsdienol. By contrast, I. pini was attracted to racemic ipsdienol, but showed no attraction to frontalin plus α‐pinene. Platysoma cylindrica was attracted to 97%‐(–)‐ipsdienol and, to a lesser extent, racemic ipsdienol, but not to frontalin plus α‐pinene. Ips grandicollis was attracted to frontalin plus α‐pinene but not to ipsdienol. Dryophthorus americanus was attracted to both ipsdienol and frontalin plus α‐pinene. 5 This ability to selectively attract the predator T. dubius without attracting the principal bark beetle in the system, I. pini, provides new opportunities for research into augmentative biological control and basic population dynamics. Moreover, the attraction of T. dubius, but not P. cylindrica, to frontalin plus α‐pinene creates opportunities for selective manipulation of just one predator. 6 Patterns of attraction by predators and bark beetles to these compounds appear to reflect various degrees of geographical and host tree overlap with several pheromone‐producing species.     

Field evaluation on the synergistic attractiveness of 2‐(1‐undecyloxy)‐1‐ethanol and ipsenol to Monochamus saltuarius

To develop an optimal attractant for Monochamus saltuarius (Gebler) (Coleoptera: Cerambycidae), the synergistic effects of a few potential attractants (ethanol and α‐pinene as host‐plant volatiles, and ipsenol and ipsdienol as bark beetle pheromones) were tested in a pine forest combined with 2‐(1‐undecyloxy)‐1‐ethanol (monochamol), the aggregation pheromone of Monochamus species, for two consecutive years, 2014 and 2015. Total number of catches was 65 and 33 in 2014 and 2015, respectively. Ethanol or ethanol + monochamol (a base blend) were not attractive to M. saltuarius with no difference from the control. Addition of α‐pinene and ipsdienol to the base blend did not significantly increase catches. However, ipsenol was significantly synergistic to the base blend in attracting M. saltuarius in 2014, and the blend (ipsenol + base blend) attracted meaningfully higher numbers of M. saltuarius in 2015. Our study illustrates the potential for monochamol and ipsenol baits for monitoring and trapping of M. saltuarius in the field.     

Aggregation pheromone of the Oriental spruce engraver Pseudips orientalis

• 1 Volatiles from the hindgut extracts of males of the Oriental spruce engraver Pseudips orientalis (Wood & Yin) (Coleoptera: Curculionidae, Scolytinae) of different phases of gallery development were analyzed by gas chromatography‐mass spectrometry‐flame ionization detection (GC‐MS/FID) with both polar and enantioselective columns.
• 2 GC‐MS/FID analyses showed that unmated males or males mated with one female produced approximately 95%‐(?)‐ipsenol and (?)‐cis‐verbenol as major components, as well as (?)‐trans‐verbenol, myrtenol, approximately 70%‐(+)‐ipsdienol and (?)‐verbenone as minor or trace components. The release of these male‐produced compounds was confirmed by GC analysis of an aeration sample of a P. orientalis‐infested spruce log. Mating reduced production of the male‐specific hindgut volatiles.
• 3 A field‐trapping bioassay in Qinghai, China, showed that a ternary blend containing two major components, 97%‐(?)‐ipsenol (i.e. close to naturally produced enantiomeric ratio) and (?)‐cis‐verbenol, plus a minor component (?)‐trans‐verbenol, caught significantly more P. orientalis beetles (♂: ♀ = 1: 2.7) compared with the unbaited control. Subtraction of (?)‐trans‐verbenol from the active ternary blend had no significant effect on trap catches. The addition of (±)‐ipsdienol (at 0.2 mg/day release) to the active ternary or binary blends significantly interrupted their trap catches. Replacing 97%‐(?)‐ipsenol with (±)‐ipsenol in the ternary blend significantly reduced trap catches to a level that was no different from the blank control.
• 4 Pseudips species were sister to all other Ipini genera in a phylogeny reconstructed with mitochondrial cytochrome oxidase I DNA data for 51 Ipini and outgroup species.
• 5 The results obtained suggest that the two major components, 95%‐(?)‐ipsenol and (?)‐cis‐verbenol (at approximately 4–5 : 1), produced by unmated fed males, are probably the primary aggregation pheromone components for P. orientalis. In light of the phylogeny, the use of terpenoid semiochemicals as pheromones probably occurred early in the evolution of Ipini and these semiochemical blends were subsequently modified in the process of speciation.

     

Geographic variation in prey preference in bark beetle predators

Abstract 1. Bark beetles and their predators are useful systems for addressing questions concerning diet breadth and prey preference in arthropod natural enemies. These predators use bark beetle pheromones to locate their prey, and the response to different pheromones is a measure of prey preference. 2. Trapping experiments were conducted to examine geographic variation in the response to prey pheromones by two bark beetle predators, Thanasimus dubius and Temnochila virescens. The experiments used pheromones for several Dendroctonus and Ips prey species (frontalin, ipsdienol, and ipsenol) and manipulated visual cues involved in prey location (black vs. white traps). The study sites included regions where the frontalin‐emitter Dendroctonus frontalis was in outbreak vs. endemic or absent. 3. There was significant geographic variation in pheromone preference for T. dubius. This predator strongly preferred a pheromone (frontalin) associated with D. frontalis at outbreak sites, while preference was more even at endemic and absent sites. No geographic variation was found in the response by T. virescens. White traps caught fewer insects than black traps for both predators, suggesting that visual cues are also important in prey location. 4. The overall pattern for T. dubius is consistent with switching or optimal foraging theory, assuming D. frontalis is a higher quality prey than Ips. The two predator species partition the prey pheromones in areas where D. frontalis is abundant, possibly to minimise competition and intraguild predation.     

Effect of different semiochemicals blends on spruce bark beetle,Ips typographus (Coleoptera: Scolytidae)

The spruce bark beetle, Ips typographus, is a recent new introduction to the Qilian Mountains of China. An outbreak of these beetles has infested over 0.03 million hectares of spruce forests in this area. Although primary attraction to volatiles has been clearly demonstrated for I. typographus, the existence and role of attraction to insect‐produced pheromones have been widely debated. Currently, commercial lures for I. typographus include only the volatiles ipsdienol, cis‐verbenol, trans‐verbenol, 2‐methyl‐3‐buten‐2‐ol and 2‐phenylethanol in Europe. Several potential pheromone candidates have been identified for I. typographus. Our GC–MS and GC–FID analyses volatiles from hindgut extracts of I. typographus in different attack phases demonstrated that the 2‐methyl‐3‐buten‐2‐ol, ipsdienol, cis‐verbenol and trans‐verbenol as major hindgut components, and ipsenol, 2‐phenylethanol, trans‐ myrtenol and verbenone as minor components. We tested various combinations of semiochemical candidates, to determine an optimal blend. Our results suggest that addition of 2‐methyl‐3‐buten‐2‐ol to either ipsenol alone, or to blends of ipsenol and other semiochemical candidates, significantly enhanced attraction of I. typographus. Therefore, a simple lure consisting of ipsenol and 2‐methyl‐3‐buten‐2‐ol would be an optimal blend of I. typographus in the Qilian Mountains, China. We conclude that this optimal semiochemical blend may provide an effective biological pest control method for use in forest ecosystem against I. typographus.     

The role of vision in the host orientation behaviour of Hylobius warreni

• 1 Visual stimuli, often in combination with olfactory stimuli, are frequently important components of host selection by forest‐dwelling phytophagous insects.
• 2 Warren root collar weevil Hylobius warreni Wood (Coleoptera: Curculionidae) is a native insect in western Canada, where larvae feed primarily on lodgepole pine Pinus contorta and can girdle and kill young trees. This weevil is an emerging problem in areas heavily impacted by mountain pine beetle Dendroctonus ponderosae Hopkins.
• 3 No olfactory attractants have been identified for this insect, making monitoring and management difficult. Thus, we investigated the role of vision in the host‐finding behaviour of Warren root collar weevil in the absence of known olfactory cues.
• 4 We conducted three experiments in field enclosure plots aiming to characterize aspects of host‐finding behaviour by adult Warren root collar weevil.
• 5 We found that both male and female weevils were readily attracted to vertical plastic silhouettes in the shape of a trunk, crown or tree at distances of less than 4 m. This pattern of attraction persisted over 2 years in two slightly different study designs. Blinding the insects removed their ability to orient to these silhouettes, indicating that host‐finding behaviour has a strong visual component. The use of different colour trunks and crowns (black, white and green) did not change the patterns of attraction of the insects to the silhouettes.
• 6 Exploiting visual attraction in this walking insect may present a new management tool in forest protection strategies.

     

Dispersal and edge behaviour of bark beetles and predators inhabiting red pine plantations

• 1 Quantifying dispersal in predator–prey systems can improve our understanding of how these species interact in space and time, as well as their relative distributions across complex landscapes.
• 2 We measured the dispersal abilities of three forest insects associated with red pine decline: the eastern five spined pine engraver Ips grandicollis (Coleoptera: Curculionidae), its main predator Thanasimus dubius (Coleoptera: Cleridae) and the basal stem and root colonizer Dendroctonus valens (Coleoptera: Curculionidae). We also examined the edge behaviours of these species and the predator Platysoma spp (Coleoptera: Histeridae) between red pine stands (habitat) and clearings (nonhabitat).
• 3 Thanasimus dubius dispersed 12 times farther than its prey I. grandicollis, with 50% of predators dispersing farther than 1.54 km. This profound difference in dispersal behaviour between prey and predator may contribute to the clumped distribution of I. grandicollis.
• 4 Most T. dubius and D. valens were confined in the pine forest, thus showing strong edge behaviour. This differed from I. grandicollis and Platysoma spp., which were commonly found in open areas adjacent to red pine plantations.
• 5 The bark beetle I. grandicollis and one of its main predators, T. dubius, exhibited different patterns of movement within a fragmented landscape. Despite a greater dispersal ability of T. dubius within forests, the spatial distribution of this predator may be restricted by fragmentation of its habitat, and provide an opportunity for partial escape of its prey.
• 6 The present study contributes to our knowledge of top‐down forces within red pine stands undergoing decline. Differences of dispersal patterns and edge behaviour could contribute to the initiation of new pockets of decline, as well as the connectedness among existing ones.

     

Density-dependent effects of multiple predators sharing a common prey in an endophytic habitat

Multiple predator species feeding on a common prey can lead to higher or lower predation than would be expected by simply combining their individual effects. Such emergent multiple predator effects may be especially prevalent if predators share feeding habitat. Despite the prevalence of endophagous insects, no studies have examined how multiple predators sharing an endophytic habitat affect prey or predator reproduction. We investigated density-dependent predation of Thanasimus dubius (Coleoptera: Cleridae) and Platysoma cylindrica (Coleoptera: Histeridae) on a bark beetle prey, Ips pini (Coleoptera: Scolytidae), in a laboratory assay. I. pini utilize aggregation pheromones to group-colonize and reproduce within the stems of conifers. T. dubius and P. cylindrica exploit these aggregation pheromones to arrive simultaneously with the herbivore. Adult T. dubius prey exophytically, while P. cylindrica adults enter and prey within the bark beetle galleries. Larvae of both predators prey endophytically. We used a multiple regression analysis, which avoids confounding predator composition with density, to examine the effects of varying predator densities alone and in combination on herbivore establishment, herbivore reproduction, and predator reproduction. Predators reduced colonization success by both sexes, and decreased I. pini reproduction on a per male and per female basis. The combined effects of these predators did not enhance or reduce prey establishment or reproduction in unexpected manners, and these predators were entirely substitutable. The herbivores net replacement rate was never reduced significantly below one at prey and predator densities emulating field conditions. Similar numbers of each predator species emerged from the logs, but predator reproduction suffered from high intraspecific interference. The net replacement rate of P. cylindrica was not affected by conspecifics or T. dubius. In contrast, the net replacement rate of T. dubius decreased with the presence of conspecifics or P. cylindrica. Combinations of both predators led to an emergent effect, a slightly increased net replacement rate of T. dubius. This may have been due to predation by larval T. dubius on pupal P. cylindrica, as P. cylindrica develops more rapidly than T. dubius within this shared habitat.     

Development of an improved attractive lure for the pine shoot beetle, Tomicus piniperda (Coleoptera: Scolytidae)

Abstract 1 The pine shoot beetle, Tomicus piniperda (L.) (Coleoptera: Scolytidae), is an exotic pest of pine, Pinus spp., and was first discovered in North America in 1992. 2 Although primary attraction to host volatiles has been clearly demonstrated for T. piniperda, the existence and role of secondary attraction to insect‐produced pheromones have been widely debated. 3 Currently, commercial lures for T. piniperda include only the host volatiles α‐pinene in North America and α‐pinene, terpinolene and (+)‐3‐carene in Europe. Several potential pheromone candidates have been identified for T. piniperda. 4 We tested various combinations of host volatiles and pheromone candidates in Michigan, U.S.A., and Ontario, Canada, to determine an optimal blend. 5 Attraction of T. piniperda was significantly increased when trans‐verbenol (95% pure, 3.2%cis‐verbenol content) was added with or without myrtenol to α‐pinene or to blends of α‐pinene and other kairomones and pheromone candidates. 6 Our results, together with other research demonstrating that trans‐verbenol is produced by T. piniperda, support the designation of trans‐verbenol as a pheromone for T. piniperda. A simple operational lure consisting of α‐pinene and trans‐verbenol is recommended for optimal attraction of T. piniperda.     

2‐(Undecyloxy)‐ethanol is a major component of the male‐produced aggregation pheromone of Monochamus sutor

The small white‐marmorated longicorn beetle, Monochamus sutor (L.) (Coleoptera: Cerambycidae), is widely distributed throughout Europe and Asia. It is a potential vector of the pine wood nematode, Bursaphelenchus xylophilus (Steiner et Buhrer) Nickle, the causal agent of the devastating pine wilt disease. Volatiles were collected from both male and female beetles after maturation feeding. In analyses of these collections using gas chromatography (GC) coupled to mass spectrometry, a single male‐specific compound was detected and identified as 2‐(undecyloxy)‐ethanol. In analyses by GC coupled to electroantennography the only consistent responses from both female and male antennae were to this compound. Trapping tests were carried out in Spain, Sweden, and China. 2‐(Undecyloxy)‐ethanol was attractive to both male and female M. sutor beetles. A blend of the bark beetle pheromones ipsenol, ipsdienol, and 2‐methyl‐3‐buten‐2‐ol was also attractive to both sexes in Spain and Sweden, and further increased the attractiveness of the 2‐(undecyloxy)‐ethanol. The host plant volatiles α‐pinene, 3‐carene, and ethanol were weakly attractive, if at all, in all three countries and did not significantly increase the attractiveness of the blend of 2‐(undecyloxy)‐ethanol and bark beetle pheromones. 2‐(Undecyloxy)‐ethanol is thus proposed to be the major, if not only, component of the male‐produced aggregation pheromone of M. sutor, and its role is discussed. This compound has been reported as a pheromone of several other Monochamus species and is another example of the parsimony that seems to exist among the pheromones of many of the Cerambycidae. Traps baited with 2‐(undecyloxy)‐ethanol and bark beetle pheromones should be useful for monitoring and control of pine wilt disease, should M. sutor be proven to be a vector of the nematode.     

Attraction to monoterpenes and beetle‐produced compounds by syntopic Ips and Dendroctonus bark beetles and their predators

• 1 Bark beetles are significant mortality agents of conifers. Four beetle species, the pine engraver Ips pini, the six‐spined pine engraver Ips calligraphus sub. ponderosae, the southern pine beetle Dendroctonus frontalis, and the western pine beetle Dendroctonus brevicomis, cohabitate pines in Arizona.
• 2 A pheromone trapping study in ponderosa forests of Arizona determined the attraction of beetles to conspecific and heterospecific pheromone components in the presence and absence of host volatiles, and tested whether predators differ in their attraction to combinations of pheromone components and tree monoterpenes.
• 3 All four bark beetle species differed in their responses to heterospecific lures and monoterpenes. Ips calligraphus was the only species that increased in trap catches when heterospecific lures were added. Heterospecific lures did not inhibit the attraction of either Dendroctonus or Ips species. The replacement of myrcene with α‐pinene increased the attraction of Dendroctonus, whereas the addition of α‐pinene had mixed results for Ips. The prominent predators Temnochila chlorodia and Enoclerus lecontei were more attracted to the I. pini lure than the D. brevicomis lure, and the combination of the two lures with α‐pinene was most attractive to both predator species.
• 4 Cross attraction and limited inhibition of bark beetles to heterospecific pheromones suggest that some of these species might use heterospecific compounds to increase successful location and colonization of trees. Predator responses to treatments suggest that tree volatiles are used to locate potential prey and predators are more responsive to Ips than to Dendroctonus pheromone components in Arizona.

     

Combining pheromone and kairomones for effective trapping of the pine sawyer beetle Monochamus galloprovincialis

Monochamus galloprovincialis (Olivier) (Coleoptera: Cerambycidae) is a secondary wood borer that acquired primordial importance since it was identified as the European vector of Bursaphelenchus xylophilus, the causal agent of pine wilt disease (PWD). An effective trapping method is needed as a tool for managing this insect vector and allowing early detection of nematode transportation. Among effective attractants identified in recent years are the specific M. galloprovincialis aggregation pheromone, host pine kairomones such as α‐pinene and bark beetle kairomones like ipsenol and methyl‐butenol. The main objective of this study was to optimize the combination of these volatiles to improve lure attractiveness and specificity. Based on ten complementary field experiments, we found a pheromone dose‐response of trap catches. The best combination of attractants was the aggregation pheromone plus two bark beetle kairomones, ipsenol and methyl‐butenol. Addition of pine terpenes, such as α‐pinene, did not significantly improve M. galloprovincialis trap capture, but did increase catch of non target species, including natural enemies. The use of pine terpenes would be advisable only if priorizing to maximize removal of vectors. While this research has lead to the development a new, highly attractive commercial lure for mature pine sawyers, none of the tested blends were successful in attracting immature pine sawyer adults. Further investigation is needed to develop attractants for these beetles.     

Complex emergence patterns in a bark beetle predator

1 The emergence pattern of Thanasimus dubius (F.) (Coleoptera: Cleridae), a common predator of the southern pine beetle, Dendroctonus frontalis Zimmermann (Coleoptera: Scolytidae), was studied under field conditions across different seasons. A simple statistical model was then developed to characterize the emergence data, using the truncated geometric distribution. Data are also presented on the mortality of T. dubius eggs at various temperatures and humidities in an effort to explain certain aspects of emergence behaviour. 2 Emergence of T. dubius from a given tree usually occurred in several discrete episodes across a two‐year period, with most individuals emerging in spring or autumn. Almost no emergence occurred in July and August, which may be an adaptation to avoid high temperature mortality. Emergence patterns appeared similar across seasons, with the time of year serving mainly to shift the pattern through time. 3 Cycles in D. frontalis abundance may be the result of delayed density dependence generated by its natural enemy complex. The predator T. dubius is likely to be an important component of this delayed density dependence, because of its lengthy development time and apparent impact on D. frontalis.     

Effects of bark beetle pheromones on the attraction of Monochamus alternatus to pine volatiles

Abstract We evaluated the attraction of Monochamus alternatus Hope (Coleoptera: Cerambycidae), Dryocoetes luteus Blandford and Orthotomicus erosus Wollaston (Coleoptera: Curculionidae: Scolytinae) to multiple-funnel traps baited with the pine volatiles, ethanol and (+)-α-pinene and the bark beetle pheromones, ipsenol and ipsdienol. M. alternatus were attracted to traps baited with ethanol and (+)-α-pinene but not those baited with ipsdienol and ipsenol. Ipsdienol and ipsenol decreased catches of M. alternatus in traps baited with ethanol and (+)-α-pinene. Traps baited with either binary combinations of ethanol and (+)-α-pinene or ipsdienol and ipsenol were attractive to D. luteus and O. erosus. The addition of ipsenol and ipsdienol to traps baited with ethanol and (+)-α-pinene synergized attraction of O. erosus but not D. luteus.     

Field response of the northern spruce bark beetle Ips duplicatus (Sahlberg) (Coleoptera: Curculionidae,Scolytinae) to different combinations of synthetic pheromone with (−)‐α‐pinene and (+)‐limonene

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Evaluation of attractants and traps for monitoring small banded pine weevil Pissodes castaneus

This study aimed to develop a semiochemical‐baited trapping system to monitor the populations of small banded pine weevil, Pissodes castaneus, a serious pest in Pinus sylvestris young stands that are weakened by biotic and abiotic factors. The scope of the work included the development of a dispenser for compounds (ethanol and α‐pinene) emitted by P. sylvestris and the pheromones of P. castaneus: grandisol and grandisal. Additionally, the effectiveness of beetle catches in different types of traps (unitrap, cross‐unitrap and long and short pipe traps) baited with a dispenser was assessed. The olfactometric studies showed that most of the newly hatched beetles that had not fed were attracted by a mixture of grandisol and grandisal. However, in the group of feeding beetles, half were attracted by a mixture of ethanol and α‐pinene. These results indicated that both pheromones and α‐pinene plus ethanol should be useful for capturing P. castaneus beetles. In the field trials, the highest efficiency was found in baited unitraps that caught up to several hundred P. castaneus beetles, while the baited cross‐unitraps caught up to a few dozen beetles. No insects were found in either type of baited pipe trap or in any of the unbaited control traps. The baited unitraps and cross‐unitraps also collected, with varied intensity, Hylobius abietis beetles, a serious pest of reforestations. These results indicate the possibility of using a unitrap baited with a 4‐component attractant for monitoring P. castaneus in integrated pest management for the protection of young forests.     

Synergistic effects of α-pinene and exo-brevicomin on pine bark beetles and associated insects in Arizona

The southern pine beetle (Dendroctonus frontalis) and western pine beetle (Dendroctonus brevicomis) cause significant mortality to pines in the southern and western United States. The effectiveness of commercial lures at capturing these bark beetles in Arizona has not been tested and may vary from other regions of their distribution. We conducted experiments using baited Lindgren funnel traps to investigate (i) if D. frontalis is more attracted to the standard commercial lure for D. brevicomis (frontalin + exo‐brevicomin + myrcene) than the D. frontalis lure (frontalin + terpene blend), (ii) whether replacement of myrcene with α‐pinene changes trap catches of Dendroctonus and associated insects, and (iii) whether the attraction to these lures varies across the geographical range of ponderosa pine forests throughout Arizona. In 2005, we tested various combinations of frontalin, exo‐brevicomin, myrcene and α‐pinene to D. frontalis, D. brevicomis and associated species. Dendroctonus frontalis, D. brevicomis and the predator Temnochila chlorodia were most attracted to lures with exo‐brevicomin. The replacement of the myrcene component with α‐pinene in the D. brevicomis lure resulted in the capture of twice as many bark beetles and Elacatis beetles. However, T. chlorodia did not differentiate between monoterpenes. In 2006, traps were set up in 11 locations around Arizona to test the relative attraction of lure combinations. In 9 out 11 locations, the D. brevicomis lure with α‐pinene was more attractive than the lure with myrcene or a terpene blend. These results suggest that the D. brevicomis lure with α‐pinene rather than myrcene is more effective lure to capture D. brevicomis and D. frontalis in Arizona. However, geographical variation in attractiveness to lures is evident even within this region of the beetles’ distributions. Differential attraction of Dendroctonus and their predators to these lures suggests potential use in field trapping and control programmes.     

Insect pheromones as reversible competitive inhibitors of acetylcholinesterase

Summary

Structural aspects of insect pheromones, especially their hydrophobicity, prompted the examination of them by in vitro and in vivo experiments as inhibitors of acetylcholinesterase. All 17 pheromones assayed in sufficient detail were reversible, competitive inhibitors of acetylcholinesterase: K_i values ranged from 0.27 mM for frontalin to 9.3 mM for ipsenol, both aggregation pheromones. All seven pheromones assayed with Tribolium castaneum immobilised this species with an effect that was weakly dose-responsive over the range of concentrations tested. Concentrations eliciting 25% immobility (IC₂₅) ranged from 2.0x10³ ppm for 3-methy 1–2-cyclohexenone to 2.0x10⁴ ppm for cis-7, cis-11-hexadecadien-1-yl acetate, both sex attractants. Such efficacies suggest that the pheromones tested would be effective as defence agents only in species living in a confined area, as under bark, or in aggregations.     

Kairomonal range of generalist predators in specialized habitats: responses to multiple phloeophagous species emitting pheromones vs. host odors

We evaluated responses of the predominant predators of pheromone-producing bark beetles (Coleoptera: Scolytidae) to chemical cues associated with other phloeophagous species that colonize the same trees. This study considered the range of chemical signals exploited by a category of predators that may be viewed either as specialists, because they feed almost exclusively within trees killed by bark beetles, or as generalists, because they feed on a diverse fauna of primary and secondary insects within this habitat. It also evaluated one aspect of a broader model of predator-prey coevolution, that proposes altered semiochemistry as a source of partial escape from predators that exploit kairomones. The predators, Thanasimus dubius (F.) (Coleoptera: Cleridae) and Platysoma cylindrica (Paykull) (Coleoptera: Histeridae), were attracted to cues associated with feeding on bark-phloem disks by two scolytids that produce adult pheromones, Ips pini (Say) and Ips grandicollis (Eichhoff). These predators were not attracted to beetles that feed on lower stems or roots and are not known to produce adult pheromones,Dendroctonus valens LeConte, Hylastes porculus Erickson (Coleoptera: Scolytidae), and Hylobius pales (Herbst) (Coleoptera: Curculionidae). The predator Tenebroides collaris (Sturm) (Coleoptera: Trogositidae) was attracted to I. pini and I. grandicollis, and also to D. valens, H. porculus, and H. pales. Ips pini was attracted to conspecifics only, but I. grandicollis was attracted both to its conspecifics and to volatiles associated with feeding lower stem and root insects. Lower stem and root insects were not or only weakly attracted to cues associated with their conspecifics. These results are consistent with a dynamic coevolved interaction between T. dubius and P. cylindrica and Ips spp.